Acta Bot. Croat. 63 (2), 147–170, 2004 CODEN: ABCRA25 ISSN 0365–0588

Naturalised species from the genus Conyza Less. (Asteraceae) in

MILENKO MILOVI]* Medical and Chemical School, Ante [upuka bb., HR-22000 [ibenik, Croatia

In this paper, the most recent state in the distribution of the American neophytes Conyza canadensis, C. bonariensis and C. sumatrensis (Asteraceae) in Croatia is presented. Conyza canadensis is believed to be one of the most widespread introduced species in the world. It arrived in the teritory of Croatia much before the beginning of the 19th century. A great number of new localities in and several new localities in the continental part of Croatia were registered in this investigation. The distribution of C. bonariensis in Croatia is limited to the Adriatic littoral, from Istria in the north to region in the south. Numerous new-discovered localities of C. sumatrensis in Dalmatia are presented. Since its first finding this neophyte has become well established and is now in the phase of expanding in the . C sumatrensis is a new species in the Croatian flora. Key words: Conyza canadensis, bonariensis, sumatrensis, flora, Croatia

Introduction The genus of Conyza Less. (Asteraceae) comprises more than 50 species, chiefly of warmer regions (MUNZ 1959). Some of them have expanded into practically all the warmer parts of the world, including Europe, mostly in man-made habitats: in the settlements, along the roads and railways and on cultivated ground. Among seven Conyza species that have been introduced into Europe so far ([IDA 2003) the most widely distributed ones are: C. canadensis (L.) Cronq., C. bonariensis (L.) Cronq. and C. sumatrensis (Retz.) E. Walker. Conyza canadensis is an annual plant believed to be a native of North America (CRON- th QUIST 1976). It was introduced into Europe in the 17 century (FOURNIER 1961; GAJI] 1975). Now it is naturalised and widespread almost throughout Europe with the exception th of the northern parts (CRONQUIST 1976). As early as the first half of the 19 century it was believed to be widely distributed in the whole area of Croatia (VISIANI 1826, 1847; PETTER 1832; ALSCHINGER 1832; SCHLOSSER and VUKOTINOVI] 1869). Conyza bonariensis (Fig. 1) is a thermo-cosmopolitan species originating from tropical America (MARTIN^I^ et al. 1999). In Europe it was introduced much later than C. cana-

* Corresponding adress: [email protected]

ACTA BOT. CROAT. 63 (2), 2004 147 MILOVI] M.

Fig. 1. Conyza bonariensis, the upper part of plant. Lateral branches overtop the main axis.

Fig. 2. Conyza sumatrensis, basal leaves in rosette. densis, and has been naturalized almost throughout the Mediterranean region and the south-western parts of Europe (CRONQUIST 1976). Findings out of this range, such as the case of an occurrence of C. bonariensis in the British Isles (STACE 1997) and the Czech Re- public (PY{EK et al. 2002; [IDA 2002; [IDA 2003) are very rare. It is established in the neigh- bouring countries of Croatia: in Italy (PIGNATTI 1982), Slovenia (MARTIN~I~ et al. 1999) and

148 ACTA BOT. CROAT. 63 (2), 2004 CONYZA IN CROATIA

Fig. 3. Conyza sumatrensis, the upper part of plant. The inflorescence is rhombic in outline.

Fig. 4. Conyza sumatrensis, detail of inflorescence.

ACTA BOT. CROAT. 63 (2), 2004 149 MILOVI] M.

Montenegro (ROHLENA 1942). The first reliable data for Croatia are given by ROHLENA (1923, as E. linifolius Willd.), for the area of Dubrovnik, and somewhat later by BOLZON (1925, as E. crispus Pourr.) for Mali Lo{inj and in the Bay of Kvarner. Since then it has been regularly noted as a naturalised species in the floras of the territory of today's Croatia (HAYEK 1931; DOMAC 1950, 1967, 1994; NIKOLI] 2000). Conyza sumatrensis (Fig. 2–4) is a neophyte originating from tropical America (PIG- NATTI 1982) and spread from there into the warmer parts of the world. The fourth volume of Flora Europaea mentions only two species of the genus Conyza (C. canadensis and C. bonariensis) but not C. sumatrensis (CRONQUIST 1976). In Europe, in France, it was introduced in the second half of the 19th century. According to ANZALONE (1964), Bonnet registered this plant in the Botanic Garden in Collioure in 1878, where it was cultivated for a long time and spread spontaneously in the town and surround- ings. At the beginning of the 20th century it was expanding throughout France, Spain and Portugal and somewhat later in Algeria in North Africa (BONNIER 1922; ANZALONE 1964). The first reliable data about the presence of this neophyte in Italy come from ANZALONE (1964, as Erigeron naudini Bonnet) who registered it in the area extending from Piemont to Sicily. Later it spread throughout the area of North Italy and for the first time it was found in the area of Trieste in 1977 (MARTINI 2000, as C. albida). The first samples of C. suma- trensis in the neighbouring Slovenia were collected by Poldini in 1984, and now it is ob- served throughout almost the whole coastal part of Slovenia (POLDINI and KALIGARI^ 2000). Recently it has been registered in Greece (DANIN 1976), Albania (BALTISBERGER and LIPPERT 1987), England (WURZELL 1988) and in Austrian Styria (MELZER 1996). In Croatia it was found for the first time in 1994 in several localities in Istria (^ARNI 1996) and later in several localities in the Bay of Kvarner (^ARNI and JOGAN 1998). But these findings were overlooked and C. sumatrensis was not registered in Flora Croatica (NIKOLI] 2000). Confusion of these three species of Conyza is possible, especially of C. bonariensis and C. sumatrensis, because of the great morphological and ecological resemblance and the lack of an appropriate literature. Such cases were reported by some authors (ANZALONE 1964; MELZER 1996; POLDINI and KALIGARI^ 2000). Identification was also made ex- tremely difficult because of the occurrence of hybrids, mostly between C. canadensis and C. bonariensis (ROHLENA 1923; HAYEK 1931; ANZALONE 1964; CRONQUIST 1976; STACE 1997). Conyza canadensis differs from C. bonariensis and C. sumatrensis by having a shorter, nearly glabrous involucre (3–4 mm long), only 25–40 female florets per capitula and hav- ing marginal florets with short (up to 1mm) but well developed ligula (CRONQUIST 1976). Conyza sumatrensis differs from closely related species C. bonariensis mostly by the marginal florets in the capitula. In the species C. sumatrensis marginal female florets are zygomorphic, while in C. bonariensis all the florets are actinomorphic. C. sumatrensis is otherwise much taller, branched out only in the upper part of the stem, lateral branches gen- erally not overtopping the main axis and the inflorescence is rhombic in outline. C. sumatrensis also has a greater number of leaves which are bigger, wider and with ramified lateral veins (PIGNATTI 1982; POLDINI and KALIGARI^, 2000; [IDA 2003). C. sumatrensis is recognisable particularly by its often well-developed winter rosettes (ANZALONE 1964, POLDINI and KALIGARI^ 2000).

150 ACTA BOT. CROAT. 63 (2), 2004 CONYZA IN CROATIA

Knowing these important diagnostic characteristics, I started my field research in the town of [ibenik and surroundings in summer 2002. In a short period of time I discovered that C. sumatrensis was established and widely distributed in this area as well as C. canadensis and C. bonariensis, but I overlooked it during my recent research (MILOVI] 2002) because I did not distinguish this plant from C. bonariensis. The field research was continued in the numerous localities in Dalmatia and also in a few localities in the conti- nental lowland part of Croatia.

Methods This investigation comprised field research as well as the analysis of the literature data and of herbarium specimens from the Croatian Herbarium (ZA) and the Herbarium of Marija and Ivo Horvat (ZAHO) in Zagreb. Field research was undertaken during the summer and autumn of 2002 and 2003. I - ited numerous localities in the area of Dalmatia, extending from Nin in the north to the Makarska region in the south, as well as Stubi~ke Toplice, Bjelovar, and Na{ice with its surroundings in the continental lowland part of Croatia. Special attention was given to investigating the distribution of C. sumatrensis as the most recent newcomer in Croatia and registered only for several localities in Istria and Bay of Kvarner. The geocoding of all localities in MTB coordinates, the new localities registered during this research as well as these identified in literature and herbaria (ZA and ZAHO), was car- ried out by the indirect method according to maps 1:100 000 (KOS et al. 2002). From among the previously registered localities, in several cases geocoding was not possible due to in- sufficiently precise citation of the location, mostly in older literature and older herbarium specimens. Specimens of Conyza species collected during this research are stored in Herbarium Croaticum in Zagreb (ZA).

Results Conyza canadensis (L.) Conq. (Erigeron canadensis L.) In the beginning of the 19th century, it was mentioned by VISIANI (1826, as Erigeron canadense) as generally and well distributed in Dalmatia. In Flora Croatica, authors already registered C. canadensis as generally and well distributed in the whole continental part of Croatia (SCHLOSSER and VUKOTINOVI] 1869, as E. canadensis L. – »In agris collibusque totius Croatiae et Slavoniae«). Since then it was registered in the major part of floristic lit- erature as well as in the literature dealing with ruderal and weed vegetation in Croatia. In Ivo and Marija Horvat Herbarium (ZAHO) there were only 4 specimens of C. canadensis: 2 from Dubravice, the district of Klanjec in Hrvatsko Zagorje, collected by Ivo Horvat and 2 from Zagreb (without precise citing of the location) collected by Brzac but defined by Horvat. In the Herbarium Croaticum (ZA) there was a total of 30 specimens collected in the area of today’s Croatia. Among them, 14 were collected in the Croatian Littoral, 12 in other parts of Croatia while 4 specimens by Vukotinovi} are without any locality cited. The old- est one is the specimen collected by Vukotinovi} in 1857.

ACTA BOT. CROAT. 63 (2), 2004 151 MILOVI] M.

The list of the previously registered localities of C. canadensis in Croatia (according herbarium and literature data analysed) are not presented in chronological order but sorted within the geographic macroregions of Croatia proposed by NJEGA^ and PEJNOVI] (2002). For these localities, geocoding was made and denotations of MTB plots (10° ´ 6°) are stated in brackets. Geocoding was impossible in these cases: no data of locality (e.g., her- barium specimens collected by Vukotinovi} in ZA); the locality is given in the broad sense (»vulgare«, »in agris et cultis totius Dalmatie frequens«, »Da«=Dalmatia), especially in older floristic works (ALSCHINGER 1832; VISIANI 1826, 1847; HAYEK 1931); and in some phytocenologic works where there are no cited locations for the individual relevés from which the synoptic (synthetic) tables of communities were made (BECK-MANNAGETTA 1901; ADAMOVI] 1911; HORVATI] 1963). Geocoding was impossible for several very small locations too, cited by ROSSI (1924) which I did not find in the maps 1:100 000 (KOS et al. 2002). These localities are presented in the following list but marked with question-marks in brackets (?). For each locality the source data are cited: author, year of publishing, for lit- erature data, as well as the author, the date of collecting and the name of the herbarium (ZA, ZAHO), for herbarium data.

The list of localities of C. canadensis registered in Croatia so far: Eastern Croatia – Slavonija, Cerevi} (?), ROSSI (Aug 1874, ZA) – around (0477,0478), HIRC (1919) – Slavonski Koba{ (0872), Slavonski Brod, Zmaj Jovina Street (0874), Velika in the Po`ega Valley (0572), Jastrebarsko (0359), Bjelovar, Kranj~evi}eva Street (0167), S. Po`ega, Panonska Street (0672), Petrinja, Rokova Street (0563), Samobor, in front of the railway station (0260), ^a|avica, near Podravska Slatina (0273); MARKOVI]- GOSPODARI] (1965) – bank of the Kara{ica River between and Satnica (0377); ILIJANI] (1967/1968) – ^a|avica near P. Slatina (0273), Osijek, the right bank of the river (0478), Velika near S. Po`ega (0572), Slavonska Po`ega, Rade Kon~ar Street (0672), Gorjani near \akovo (0676), Slavonski Koba{ (0872), Slavonski Brod, Poloj (0874), @upanja (0978); MARKOVI] (1975, 1979, 1981) – ^rnkovci (0275), Petrijevci (0377), Osijek, the road to Darda (0478); TOPI] (1978) – Dra` (0178), Bocanjevci (0375), Vinogradci (0376), Petrijevci (0377), Valpova~ka Gorica (0476), Beli{}e, along the Drava river (0376), Zel~in near (0376), ^epinski Martinci (0476), Vukovar, Vukovarska ada (0680), Borovo, Borovska ada (0679), (0776), \ur|anci (0777), Divo{evci (0876); RAU[ and [EGULJA (1983, 1985); [EGULJA and TOPI] (1987) – Tenja (0478,0578); PAN^I] (1984) – Po`ega valley, village of Nurkovac near Po`ega (0671), TOMA[EVI] (1972, 1998) – settlements in the region of \akovo (0676,0776), MARKOVI] (1990)

Central Croatia – Vara`din, Slavenska Street (9664), Samoborsko gorje, Kon{}ica (0260), Remetinec (9863) and Klju~ (9864) near Novi Marof; MARKOVI]-GOSPODARI] (1965) – Mursko Sredi{}e (9464) and Kri`ovljan (9765) in the surroundings of Vara`din, Jesenice near Zagreb (0160); MARKOVI] (1984a)

152 ACTA BOT. CROAT. 63 (2), 2004 CONYZA IN CROATIA

– Krapinske Toplice (9961); [O[TARI] and MARKOVI] (1998) – Dubravica in the district of Klanjec (9960), HORVAT (Aug 1918 and Aug 1950, ZAHO) – Ivan{~ica in Hrvatsko Zagorje (9862), MARKOVI] and REGULA-BEVILACQUA (1988) – Strahin{~ica Mt. in Hrvatsko Zagorje: @utnica (9861), Prese~ina stream (9861), the north side of Strahin{~ica (9861), Radoboj near village of Tu{ki (9861), Donje Stra- hinje (9861); REGULA-BEVILACQUA (1979); REGULA-BEVILACQUA and [EGULJA (2000) – Konj{~ina, Turni{}e (9962), STAN^I] (1994) – Kri`evci, Maren~i}eva Street (9965), ^azma (0265), Vara`din, railway bridge over the Drava River (9664), Zagreb, Savski most (0261); MARKOVI] (1975) – area of \ur|eva~ki pijesci (9968), Molve (9868), Vukosavljevica (0169); SOKLI] (1943) – Koprivni~ki Ivanec and Pustakovec (9766), \elekovec and Mali otok (9766,9767), Mo~ilski Breg (9866), Peteranec and Sigetec (9867), Ivane~ki Botinec and Virje (9967,9968); KOVA^EVI] (1957) – Zagreb (?); ROSSI (Sept 1883, ZA), BRZAC (Sept 1948, ZAHO, 2 sheets) – Zagreb, railway bridge over the River and Savska Street (0261); HORVATI] (1947); GOSPODARI] (Aug 1954, ZA) – Zagreb: Podsused and Jarun (0261), @itnjak and Petru{evac (0262); HORVATI] and GOSPODARI] (1959/1960); MARKOVI] (1973, 1978) – Medvednica, departure station of the cable railway (0161), MARKOVI] (Aug 1960, ZA); Medvednica, area of Drage in Vugrovec (0162), HULINA (1979); Zagreb, Slobo{tina (0262), KUMBARI] (Jul 1992, ZA) – Zagreb, Podsused (0161), Ivanja Rijeka near Zagreb (0262), Galdovo Erdotsko near Sisak (0564); MARKOVI] (1980) – Turopolje: Lukavac (0261), Hra{}e (0262), Dragono`ec (0361), Gudci (0361), Obre` (0362), Petrovina (0362), Vukovina (0362); HULINA (1989, 1991); GRA^ANIN et al. (1971) – Zden~ina, {uma »[ikara« (9864), [TEFANAC (1957, ZA); Crna Mlaka (0360), [TEFANAC (Aug 1957, ZA); Vukomeri~ke Gorice (0361), [EGULJA (1977) – Kostajnica (0765) and Maja (0762); [EGULJA et al. (1998) – Karlovac and the surroundings: small forest of Lu{~i} (?), Logori{}e (0559), Mahi~no (0559); ROSSI (Aug 1886, Sept 1908 and Aug 1918, ZA); ILIJANI] (1961/1962) – Samobor and Gornji Kraj (0260); Banovina region: Hrastovica (0563,0663), (0662), Topusko and Vranovina (0761); wider area of Karlovac and Slunj: Ribnik and Stative (0458), Orlovac (0459), Gradec (0461), Mrzlopolje (0558), Selce (?), Banija (?), the forest of Debela Glava (0558), Dubovac, Turanj and Vukmani} (0559), small forest of Lu{~i} (?), Rakovac (?), Krnjak (0659), Vojni} (0660), Slunj and Slunjska Brda (0859); ROSSI (1924, 1930), GJURA[IN (Aug 1889, ZA)

Mountain Croatia – Gorski Kotar (?), MARKOVI] (1984b) – Ogulin and Sovinica (0757), hill of Mo{unje (?), Dre`nica (0856), Znidar{i} (?), Modru{e (?), Jezerane (0957), Vratnik and @uta Lokva (1056), Bro}anac and Rakovica (1059), Lake of Milanovac (1159); ROSSI (1924, 1930) – Plitvice, Lake of Milanovac (1159); ROSSI (Jul 1911, ZA) – Kapela Koreni~ka (1160) and Kapela (1260); [EGULJA (2000)

ACTA BOT. CROAT. 63 (2), 2004 153 MILOVI] M.

– Sv. Kri` (1055), Vratnik and Senjska Draga (1056), Gaj near Pisarola (?), Takalice near O{tarije (0757), Prozor near Oto~ac (1157), Gospi} (1458), [tikada (1660), Ri~ice (1660), Gra~ac (1761), near @egar (1861); DEGEN (1938)

North Croatian Littoral – Rovinj, St. Andrija and St. Katarina (0947); BENACHIO (1939) – Vodnjan (1049), Gali`ana (1048), Fa`ana (1048), Pula (1149), Veruda (1149), Preman- tura (1249); FREYN (1877) – wider area of Rt Kamenjak (1249); TOPI] and [EGULJA (2000) – Pore~ (0757), Ra{a (0950), Labin (0950); ^ARNI (1996) – NE part of the Labin area (0950); [EGULJA (1981) – Rijeka (0652), Bakar (0653), Kraljevica (0753); HIRC (1884); ROSSI (1924, 1930) – Rijeka (0652), Islands of the Bay of Kvarner (?); HAYEK (1931) – Rijeka, Marx and Engels av., Nr 9 (0652), Senj (1055); MARKOVI] (1964) – [krljevo near Rijeka (0653); RANDI] (Aug 1981, ZA) – Kraljevica, near the Veritas restaurant (0753), Rijeka, in the port (0652), Crikvenica (0854); ^ARNI and JOGAN (1998) – Senj (1055); DEGEN (1938); MARKOVI] (Aug 1962, ZA) – Island of Krk, »frequente in tutta l’is. di Veglia«, (0753,0852,0853,0952,0953,0954, 1054); LUSINA (1932, 1934) – Island of : Cres (1052), NW part of Vransko jezero (1152); HIRC (1913); HIRC (Sept 1903, ZA); MARKOVI] (Aug 1962, ZA, 2 sheets); MARTINI (1990) – Island of (1452), Island of , the settlement of Unije and Campo (1351); LUSINA (1936, 1956) – Mali Lo{inj: ^igal (1452), Sv. Jakov (1352), the Bay of Kovcanja (1452); HARA^I] (Oct 1889, Oct 1890 and Aug 1899, ZA) – Island of (1451); HARA^I] (Jul 1890, ZA); LUSINA (1933); HODAK (1956)

South Croatian Littoral (Dalmatia) – (1857), ALSCHINGER (1832) – (?); PLAV[I]-GOJKOVI] (1966), TRINAJSTI] (1991) – National Park: near village of Rupe, Dubravice and area between Skradin and Bi}ine (2161); MARKOVI] et al.(1993) – Betina on the Island of (2159), the Island of (2261, 2361), the Island of (2361), Prvi} Island (2260), small islands near Murter: Pri{njak, Zminjak and Veliki Vinik (2159); PAND@A (1998a, 1998b, 1998c, 2002) – [ibenik: Varo{ and Crnica (2261); MILOVI] (Sept 1982, Aug 1996, ZA) – Pirovac (2160), [ibenik (2261), Bilice, Novo naselje (2261), Lozovac (2261), Rupi}i (2261), Dubrava, [kugori and [i{ak (2261), Jadrija (2261), Zabla}e (2261), Konje- vrate, Krni}i (2262), Danilo Gornje, Bedrice and Grande{i (2262), Kraljice (2262), Solaris (2361), Jadrtovac (2361), Greba{tica (2361), Perkovi} (2362), Boraja, Kne`e- vi}i (2362); MILOVI] and RANDI] (2001); MILOVI] (2002) – Mt Mosor, Gornje Sitno (2465); BEDALOV and [EGULJA (1987) – The Bay of Ka{tela (2464); [OLI] et al. (1998) – Split (2464); PETTER (1832), RU[^I] (2003)

154 ACTA BOT. CROAT. 63 (2), 2004 CONYZA IN CROATIA

– The Island of Veli Drvenik (2562), BEDALOV (1976); The Island of Mali Drvenik (2562), VLADOVI] et al. (2002) – The Island of (?); TRINAJSTI] (1993) – The Island of Bra~: Milna (2664), Brdo and Bobovi{}e (2664); BELLOTI (Aug 1976, ZA); [TAMOL and MARKOVI] (1985) – The island of [olta (2663); BEDALOV (1989) – Makarska coastal region (2667,2668,2768); [ILI] and [OLI] (1999) – The Island of (3267); TRINAJSTI] (1979) – The Island of Vis, throughout the Island (2962,2963); DOMAC (1955) – The Island of (2960); PAVLETI] (1980) – The Island of Bi{evo (3062); GINZBERGER (1921) – Mali Ston (3172), Trstenik and Dra~e (3070) in the peninsula of Pelje{ac; JASPRICA and KOVA^I] (1997a, 1997b); TRINAJSTI] and JASPRICA (1998) – The Island of , Pinjevica (3272); REGULA-BEVILACQUA and JURKOVI]-BEVILAC- QUA (1980) – Sr| and the surroundings of Dubrova~ka rijeka (3374); BIRA^ (1973) – The Island of Badija, in the village of Badija (3069); BAR^I] (1974); TRINAJSTI] (1985, 1995) – The Island of [ipan: [ipanska Luka and Su|ura| (3273); HE]IMOVI], M. (Aug 1979, ZA); HE]IMOVI], M. (1981, 1984) – The Island of Lokrum, Botanic Garden (3374); HE]IMOVI], S. (Aug 1979, ZA); HE]I- MOVI], S. (1982) – The Island of (3373); the Island of Kolo~ep, Gornje and Donje ^elo (3374); HE]IMOVI],M.andHE]IMOVI], S. (1986, 1987)

The list of new localities registered during this research: – Stubi~ke Toplice (0061); widespread in the settlement and surroundings; herb., 30 Aug 2002, near Matija Gubec Hotel – Malo Korenovo near Bjelovar (0166); widespread in the settlement and surroundings; herb., 16 Jul 2002 – Narta near Bjelovar (0166); widespread in the settlement; obs., 16 Jul 2002 – Valpovo (0376); widespread in the settlement and surroundings; herb., 11 Jul 2003, in Kolodvorska Street – Ko{ka (0475), (0476), Markovac Na{i~ki (0574), Na{ice and Gradac Na{i~ki (0574), ^aglin and Ru{evo (0673), Podcerkavlje near S. Brod (0774); very common along the streets in settlements; obs. 15 Aug 2003 – Benkovac (1559); widespread: Primary School, Market place, railway and bus station; herb., 1 Nov 2002, railway station – Nin (1757), @drijac and Zukve (1757); widespread on ruderal sites; obs., 20 Jul 2003. – Obrovac (1760), Bibinje (1957), Suko{an (1957); widespread throughout the settle- ments; obs., 1 Nov 2002 – Knin (1963); widespread in the town and its surroundings; obs., 30 Aug and 16 Nov 2002.; herb., 16 Nov 2002, near railway station – Turanj and Filip Jakov (2058), Biograd (2058), Pako{tane (2059); common by roads and in gardens; obs., 17 Nov 2002 – Stankovci (2060); by the road through the settlement; obs., 1 Nov 2002

ACTA BOT. CROAT. 63 (2), 2004 155 MILOVI] M.

– Drni{ (2162); widespread in the settlement and surroundings; herb. 16 Nov 2002, near the church »Gospa od Ro`arije« – Tribunj (2260); by roads near the coast; herb., 20 Oct 2002, by the road in the settlement – Vodice (2260); common in some places in the town and surroundings; herb., 20 Oct 2002, by the road near the Punta Hotel – Srima (2260); by the roads and as a weed in gardens; obs., 20 Jul 2003 – Une{i} (2263); very common in the settlement and its surroundings; obs., 14 Oct 2001. – The Island of @irje (2359,2360): Muna, @irje, @irjansko polje; common in the ruderal sites in the settlement and as a weed in vineyards and fields; obs., 27 Oct 2001 – The Island of Obonjan (2360); rare, probably came recently; herb., 24 Aug 2002, in the ruderal vegetation near swimming pool and restaurant – Brodarica (2361); very common in the settlement; obs., 17 Sept 2001 – Primo{ten (2461); common in the surroundings of the Bay of Porat; obs., 16 Dec 2002 – Rogoznica (2461), very common in the whole settlement; obs., 16 Dec 2002 – Trogir (2463); in ruderal vegetation by the streets near the market; obs., 19 Oct 2002

Fig. 5. Geographic distribution of C. canadensis in Croatia. previously registered localities, · new localities

156 ACTA BOT. CROAT. 63 (2), 2004 CONYZA IN CROATIA

Conyza bonariensis (L.) Cronq. Synonyms: Conyza ambiqua DC., Erigeron crispus Pourr., E. linifolius Willd. According to the available literature, the first reliable data concerning the establishment of C. bonariensis in Croatia were from ROHLENA (1923, as E. linifolius Willd) for the area of Dubrovnik (»circa Dubrovnik, praecipue in peninsula Lapad frequens«). Somewhat later it was found by BOLZON (1925) in Mali Lo{inj and Krk in the Bay of Kvarner. In the first half of the 20th century it was registered also for the islands of Kvarner (Lo{inj, Unije and Male Srakane) by LUSINA (1936) and for the Island of Kor~ula in 1940 by MAILLEFER (according to TRINAJSTI] 1985). Not a single specimen of this neophyte was stored in ZAHO while in ZA there were only 6 specimens, collected in the area of Croatian Littoral in the period from 1961 to 1996. Because of the possibility of confusion with C. sumatren- sis, all the specimens of C. bonariensis were carefully examined but they turned to be cor- rectly defined.

The list of previously registered localities of C. bonariensis in Croatia:

– The Island of Krk, Krk (0953); the Island of Lo{inj, Mali Lo{inj (1452); BOLZON (1925) – Rovinj (0947), Krk (0953); SCHULTE (1989) – The Island of Susak (1451); HODAK (1956) – The Island of Lo{inj: wider area of Mali Lo{inj (»Cigale, Calvario, Duomo, Istmo, Curila, Poliana«) (1452); the Island of Unije, settlement and Campo (1351); the Island of Male Srakane (1452); LUSINA (1936); MARKOVI] (Aug 1962, ZA) – The : Rupe (2161), Dubravice (2161), Skradin (2161), Raslina (2261), Mikulandre and Vrulje (2261); MARKOVI] et al. (1993) – [ibenik and surroundings: [ibenik (2261), Jadrija (2261), Zabla}e (2261), Solaris (2361), Jadrtovac (2361), Greba{tica (2361), Bilice (2261), Konjevrate (2262), Rupi}i (2261), Dubrava (2261), Vrpolje (2362), Perkovi} (2362), Boraja (2362); MARKOVI] (1964); MILOVI] (Jun 1996, ZA); MILOVI] (2002) – The Island of Murter: Murter (2159), Betina (2159), Tisno (2259), Jezera (2259); small islands of Murter: Pri{njak, Tegina, Veliki Vinik and Sustipanac (2159); PAND@A (1998a, 2002) – The Island of Zlarin (2261, 2361), the Island of Prvi} (2260), the Island of Krapanj (2361), the Island of (2360); FRANJI] and PAND@A (1996); PAND@A (1998b, 1998c) – The Island of @ut (2157); GA@I-BASKOVA and BEDALOV (1978) – The Bay of Ka{tela (2464); [OLI] et al. (1998) – Split (2464); RU[^I] (2003) – The Island of ^iovo (2463), the Island of Bra~, Sutivan (2664), the Island of Kor~ula, Lumbarda (3069); MARKOVI] (1964) – The Island of Bra~: Bijaka, Vla{ka, Raci} and Brdo (2664), road from Milna to Bobovi{}e (2664), Sutivan (2664); BELLOTTI (Jun 1976, ZA); [TAMOL and MARKOVI] (1985) – Makarska coastal region (2667,2668,2768); [ILI] and [OLI] (1999) – The Islands of Palagru`a (3663), the Island of Svetac (2960); PAVLETI] (1978, 1980) – The Island of Kor~ula (?); MAILLEFER 1940 (according TRINAJSTI] 1985)

ACTA BOT. CROAT. 63 (2), 2004 157 MILOVI] M.

– The Bay of Mali Ston (3172); Peninsula of Pelje{ac: Trstenik (3070), Dra~e (3070), Janjina (3070); JASPRICA and KOVA^I] (1997a, 1997b) – The Island of [ipan (3273); HE]IMOVI], M. (Aug 1979, ZA); HE]IMOVI], M. (1981) – The Island of Lokrum, Botanic Garden (3374); HE]IMOVI], S. (Jun 1979, ZA); HE]I- MOVI], S. (1982) – The Island of Lopud (3373); the Island of Kolo~ep, Gornje and Donje ^elo (3374); HE]IMOVI], M. and HE]IMOVI], S. (1986, 1987) – Dubrovnik, the peninsula of Lapad and around the town (3374); ROHLENA (1923) – Cavtat, in the port (3475); MARKOVI] (May 1961, ZA)

The list of new localities of C. bonariensis: – Benkovac (1559); rare, near the market place, on the railway station; herb., 1 Nov 2002, by the street near market place – Zadar, Vo{tarnica(1857); in the ruderal vegetation beside roads and pathways; herb., 17 Nov 2002, Vo{tarnica, by the road – Bibinje (1957); in the ruderal vegetation beside roads and pathways, trampled sites; obs., 17 Nov 2002 – Suko{an (1957), widespread in ruderal vegetation beside roads and pathways in the set- tlement; obs., 17 Nov 2002 – Turanj and Filip Jakov (2058), common beside roads and pathways; obs., 17 Nov 2002. – Biograd (2058); common on ruderal sites in the settlement; herb., 17 Nov 2002, grass- land by the road, between the School and clinic – Pako{tane (2059); very common in the ruderal sites in the settlement; herb., 17 Nov 2002, trampled site by the street near the Adriatic autocamp – Pirovac (2160); common in the ruderal sites, grasslands and gardens in settlement; herb., 20 Oct 2002, untilled garden near Pirov~anka department store – Tribunj (2260); widespread in the settlement; herb., 20 Oct 2002, by road near coast – Vodice (2260); common by the roads and in the cracks of pavements; herb., 20 Oct 2002, grassland near Punta Hotel – Srima (2260); common in settlements, on trampled habitats by roads and pathways; obs., 20 Sept 2003 – The Island of @irje: Muna, @irje (2359), @irjansko polje (2360); ruderal habitats in the settlement, in vineyards, untilled fields; obs., 27 Oct 2001 – The Island of Obonjan (2360); rare, on ruderal sites; herb., 24 Aug 2002, ruderal vege- tation near swimming pool and restaurant – Brodarica (2361); widespread by the roads and pathways, on grasslands; obs., 18 Oct 2002 and 26 Jul 2003 – Primo{ten (2461); ruderal sites in the settlement and surroundings; herb., 16 Dec 2002, Porat, untilled vegetable garden – Rogoznica (2461); widespread in the settlement and surroundings; herb., 16 Dec 2002, by the road near Frappa Marina – Trogir (2463); common by the streets, pathways, grasslands and gardens; obs., 23 Nov 2002

158 ACTA BOT. CROAT. 63 (2), 2004 CONYZA IN CROATIA

Fig. 6. Geographic distribution of C. bonariensis in Croatia. previously registered localities, · new localities

Conyza sumatrensis (Retz.) E. Walker Synonyms: Conyza albida Willd. ex Sprengel, C. altissima Naud. et Deb., C. naudini Bonet Not a single specimen of C. sumatrensis was stored in the two largest public herbaria in Croatia (ZA, ZAHO). This neophyte was registered for the first time in Croatia in summer 1994 in Istria within thermophilous vegetation of trampled habitats classified into the Euphorbion prostratae alliance, the Eragrostietalia,theStellarietea mediae (^ARNI 1996, as C. albida), on these localities: – Pore~, in front of the Obala restaurant (0747) – Vrsar, in the town (0847) – Rovinj, in front of the Adriatic restaurant and on the coast near the shipyard (0947) – Pula, on the coast, in front of the department store, in front of the military Command, in the port, in front of the Uljanik shipyard and at the railway station (1149)

ACTA BOT. CROAT. 63 (2), 2004 159 MILOVI] M.

– Rabac, along the road on the coast (0950,0951) Soon after that it was registered also on the several localities in the Bay of Kvarner within similar type of vegetation (^ARNI and JOGAN 1998, as C. albida) – Lovran, in front of the hotel Excelsior (0751) – I~i}i, in the marina (0651) – Opatija, Viktora cara Emina Street (0651) – Volosko, in the street above the church (0651) – Rijeka, Trsat, in front of the church (0652) – Bakar, on the sand at the shore (0653) – Kraljevica, along the street near the Veritas restaurant (0753) – The Island of Krk: Vrbnik, in front of the church and Vitezovi}eva Street (0954); Krk, between the port and bus station (0953); Ba{ka, along the path (1054); Punat, Hotel Park parking lot (0953) – Senj, Pavlinski trg (1055) – The Island of : Rab, the church of St. Euphemia (1254) – The Island of Lo{inj: Mali Lo{inj, on the sand at the shore and in front of the local church (1452)

The list of new localities of C. sumatrensis: – Obrovac (1760); rare, probably common in the recent time; herb., 1 Nov 2002, ne- glected flower garden near the School – Benkovac (1559); common on ruderal sites; herb., 1 Nov 2002, grassland near market – Nin, old part of town (1757), @drijac and Zukve (1757); common by roads and path- ways in settlements; obs., 20 Jul 2003 – Zadar (1857); widespread in ruderal sites in the town and on the fields in surroundings; herb., 17 Nov 2002, Vo{tarnica, by the road – Bibinje (1957); widespread in the settlement and surroundings; herb., 17 Nov 2002, [pirini Street, by the road near the railway – Suko{an (1957); widespread in the settlement by the roads; obs., 17 Nov 2002 – Turanj and Filip Jakov (2058); common by the roads and pathways near coast; obs., 17 Nov 2002 – Biograd (2058); common on ruderal sites in the town; herb., 17 Nov 2002, ruderal veg- etation between the School and clinic – Pako{tane (2059); widespread in the whole settlement; herb., 17 Nov 2002, by the road near the Adriatic autocamp – Stankovci (2060); common by the main road through settlement; obs. 1 Nov 2002 – Pirovac (2160); widespread by the roads, on grasslands and gardens; herb. 20 Oct 2002, neglected garden near the Pirov~anka department store – The Island of Murter: Betina and Murter (2159); Tisno and Jezera (2259); ruderal sites in settlements, gardens, untilled fields in surroundings; obs. 1 Nov 2002. and 4 Jul 2003 – Tribunj (2260); very common by the roads and pathways near the coast, trampled sites; herb., 20 Oct 2002., by road near coast – Vodice (2260); common by the roads and pathways; herb., 20 Oct 2002, by the road near the Punta Hotel – Srima (2260); common by the roads; obs., 20 Oct 2002

160 ACTA BOT. CROAT. 63 (2), 2004 CONYZA IN CROATIA

– The Island of Prvi} (2260): widespread in the settlement of Luka and [epurine, by roads, in grasslands and gardens; herb., 26 Oct 2002, Prvi} Luka, untilled vegetable garden – [ibenik (2261); widespread in the whole town and its surroundings; herb., 13 Aug 2002, Vidici, by the road; 19 Aug 2002, [ubi}evac, on the bottom of the wall – Zabla}e (2261); common in the settlement; herb. 16 Dec 2002., trampled grassland by the road in the central part of settlement – Solaris (2361); very common in ruderal sites through tourist complex; herb., 4 Aug 1996, mistaken for C. bonariensis – The Island of Zlarin (2261,2361), widespread in settlement and Zlarinsko polje; herb., 23 Aug 2002, by road in village – Brodarica (2361); widespread in the settlement and surroundings, by the roads, path- ways, trampled grasslands, neglected gardens; obs. 18 Jun 2002 and 26 Jul 2003 – The Island of Krapanj (2361); widespread on the whole Island; herb., 18 May 2002, by road near the coast – The Island of @irje: Muna and @irje (2359), @irjansko polje (2359,2360); common in the settlement and in the area of Zlarinsko polje; obs., 27 Oct 2001 – The Island of Obonjan (2360); rare, probably approached recently; herb., 24 Aug 2002 on ruderal site near swimming pool and restaurant – Skradin, (2161); widespread in town and surroundings; herb., 5 Nov 2002, by the road near the football stadium – Drni{ (2162); only few specimens in front of church Gospa od Ro`arije, probably came recently; herb., 16 Nov 2002 – Une{i} (2263); rare in the settlement and surroundings; obs., 16 Nov 2002 – Perkovi} (2362); widespread in the whole settlement, mostly in the area of railway sta- tion; obs. 16 Nov 2002 – Vrpolje (2362); widespread by the roads in the settlement and on fields in surroundings; obs., 16 Nov 2002 – Greba{tica (2361); widespread through the whole settlement; herb. 16 Dec 2002, by the road along the coast – Primo{ten (2461); common in the whole settlement, by the roads and pathways; herb. 16 Dec 2002, untilled vegetable garden by small Bay of Porat – Rogoznica (2461); widespread in the whole settlement; herb. 16 Dec 2002, by the road near Frappa Marina – Trogir (2463); by the roads and pathways near bus station and market; obs., 23 Nov 2002 – Split (2464); widespread in the whole town and its surroundings; herb., by the road near railway station, 23 Nov 2002 – Jesenice (2565), by the roads; Omi{ (2566), common on ruderal sites in town; obs., 19 Oct 2002 – Omi{, Radmanove mlinice on the River (2566); rare; herb., 19 Oct 2002, by the road – Makarska (2668,2768), widespread on ruderal sites in the whole town and its surround- ings: Brela, Ba{ka Voda and Krvavice (2667); herb., 19 Oct 2002, Makarska, by the road along the coast near the Biokovka Hotel

ACTA BOT. CROAT. 63 (2), 2004 161 MILOVI] M.

Fig. 7. Geographic distribution of C. sumatrensis in Croatia. previously registered localities, · new localities

Discussion Among the three Conyza species investigated, C. canadensis became established in Croatia a long time ago. In the botanic literature from the 19th century already, this annual neophyte was recognized as a widespread and naturalized species in the Croatian littoral (VISIANI 1826, 1847; FREYN 1877) as well as in the continental lowland parts of Croatia (SCHLOSSER and VUKOTINOVI] 1869). The results of the analysis of numerous botanic works in the 20th century and the analysis of herbarium specimens (ZA, ZAHO) as well as the great number of new localities registered during this research confirm that this species is one of the most common adventitious plants in the appropriate habitats in the whole of Croatia (Fig. 5). Conyza canadensis most frequently occurs within nitrophilous weed comunities on cultivated land and ruderal communities in the settlements classified within the Chenopo- dietalia (the Chenopodietea) in the coastal region of Croatia (HORVATI] 1963) as well as in its continental regions (MARKOVI]-GOSPODARI] 1965; RAU[ and [EGULJA 1985). In the

162 ACTA BOT. CROAT. 63 (2), 2004 CONYZA IN CROATIA continental, lowland parts of Croatia it frequently occurs in the nitrophilous ass. Polygono- -Bidentetum (the Bidentalia tripartiti, Bidentetea tripartiti) which grows on natural and seminatural habitats along river banks but also in antrophogenic habitats in villages: along channels, ditches, swamps and pasture-grounds (MARKOVI] 1975; RAU[ and [EGULJA 1985). In Istria and the Bay of Kvarner it occurs in the thermophilous vegetation of tram- pled habitats from the Eragrostietalia,theStellarietea mediae (^ARNI 1996; ^ARNI and JOGAN 1998). Conyza bonariensis was not mentioned in the classic floristic works of the 19th century concerning the territory of today’s Croatia (VISIANI 1826; SCHLOSSER and VUKOTINOVI] 1869; FREYN 1877) but was registered for the first time for the area of Dubrovnik (ROHLENA 1923) and after that on the Kvarnerian islands: Krk, Lo{inj, Unije and Male Srakane (BOLZON 1925; LUSINA 1936). The analysis of the literature and herbarium data show that majority of the findings were from the second part of the 20thcentury. This analysis indi- cates that C. bonariensis was established in Croatia in the first half but spread along the Croatian littoral in the second half of the 20th century. There have been no data of findings of this species in the continental part of Croatia so far. Numerous new localities in the area of Dalmatia, from Nin in the north to Makarska in the south, were registered during this re- search. With exception of Benkovac, C. bonariensis was not found in the settlements of the Dalmatian hinterland (Obrovac, Knin, Drni{ and Une{i}) researched. It occurs frequently in communities on the ruderal habitats (the Erigero-Xanthietum and Hordeetum leporini) and also in communities in cultivated soil (the Tribulo-Amaranthetum and Fumario-Cype- retum rotundi) from the Chenopodietalia,theChenopodietea (HORVATI] 1963; MARKOVI] 1964; HE]IMOVI], M. 1981). The range of C. bonariensis in Croatia so far (Fig. 6) has been limited to the narrow area of the Adriatic coast with the islands, extending from Istria in the north to the region of Dubrovnik in the south. The majority of localities are situated in the Mediterranean zone proper with potential natural vegetation from the Quercion ilicis alliance. This tropi- cal-subtropical neophyte has become well acclimatized and widely spread along the Cro- atian littoral with a dry and warm Mediterranean climate. Conyza sumatrensis, origin from tropical America, was introduced in Europe in the sec- th ond part of the 19 century (Bonnier 1878, according to ANZALONE 1964). Since then it has been expanding mostly into the warmer parts of the Mediterranean region. According to the literature data the most invasive phase has been since the middle of the 20th century so far (ANZALONE 1964; DANIN 1976; BALTISBERGER and LIPPERT 1987; WURZELL 1988; MELZER 1996; ^ARNI 1996; POLDINI and KALIGARI^ 2000). Among the three researched species of Conyza, C. sumatrensis was established in Croatia the latest. The first and only findings for Croatia were registered in Istria and the Bay of Kvarner where it occurs in thermophilous vegetation on trampled sites from the Eragrostietalia,theStellarietea mediae (^ARNI 1996; ^ARNI and JOGAN 1998). These find- ings were overlooked and this plant was not mentioned in Index of Flora Croatica by NIKOLI] (2000). During this research C. sumatrensis was found in numerous new localities in Dalmatia, from Nin in the north to Makarska in the south (Fig. 7). In several of the most recent investi- gations of the flora of Dalmatia, authors have not mentioned this species (FRANJI] and PAND@A 1996; PAND@A 1998a,b,c; [ILI] and [OLI] 1999) probably because they have not

ACTA BOT. CROAT. 63 (2), 2004 163 MILOVI] M. distinguished it from C. bonariensis. This was the case during my most recent research into the flora of the wider area of [ibenik (MILOVI] and RANDI] 2001, MILOVI] 2002). The con- fusion of C. sumatrensis for C. bonariensis also happened in the neighbouring countries of Austria and Slovenia (MELZER 1996; POLDINI and KALIGARI^ 2000). The geographic distribution of C. sumatrensis so far has been limited to the coastal re- gion of Croatia: Istria, the Bay of Kvarner and Dalmatia, extending to the Makarska region in the south (Fig. 7). It occurs in dense populations in a large number of localities in Dalmatia, especially in the wider area of [ibenik. In the last ten years, it has become well acclimatized almost throughout the Croatian coast, which is also due to the warm Mediter- ranean climate of this area. We can suppose it was already spread in the rest of Croatian lit- toral, south of Makaraska region as well as on some of the islands. Through their native geographic range all the three species of Conyza occur in open sites in abandoned fields, vineyards, waste areas, railways and roadsides, and disturbed sites in natural communities (MUNZ 1959; THEBAUD and ABBOTT 1995). It occurs in the Croatian littoral on the similar habitats too, but more frequently in ruderal sites in the settle- ments and its surroundings than in cultivated soil. Following the recent proposal of »invasiveness« given by RICHARDSON et al. (2000), all the three Conyza species are considered invasive in Spain (SANZ-ELORZA et al. 2001; DANA et al. 2001). C. canadensis was the one of the most common and widespread weed (LORENZI and JEFFREY 1987). As a weed it was considered invasive in the region of former Yugoslavia and today’s Croatia, too ([ABI] ed. 1982; KOJI] 1985; HULINA 1991,1998). In recent times C. bonariensis and C. sumatrensis have become, widespread and dangerous weeds in agricultural and horticultural practice and in the invasion of natural and semi- natural vegetation (THEBAUD and ABBOTT 1995; PRATHER 2000; SANZ-ELORZA et al. 2001). The expansion of C. bonariensis and C. sumatrensis has been limited to the coastal re- gion of Croatia so far, for these plants prefer the warm climate of the Mediterranean proper without or with only late frost. It can be expected not to expand out of this area in future, but individual findings cannot be excluded. Such cases are seen in the sporadic occurrence of these species out of the Mediterranean region: C. sumatrensis in Austrian Styria (MELZER 1996), C. bonariensis in the Czech Republic (PY[EK et al. 2002; [IDA 2002, 2003) and both of the species in the British Isles (STACE 1997). The occurrence of these tropical-subtropi- cal species out of the warm regions especially in the big cities (London) can be explained as a consequence of »global warming« and of the phenomena grouped as »heat island effect«.

Acknowledgements I am grateful to Dr Toni Nikoli} who prepared the maps, as well as to the anonymous re- viewer and the editor Damir Vili~i} for critical comments on the previous version of the manuscript. Thanks also go to my colleague Mislava Dragutin for language assistance in writing this article.

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